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Revision of the genus Aseptis McDunnough
(Lepidoptera, Noctuidae, Noctuinae, Xylenini) with a
description of two new genera, Paraseptis and Viridiseptis

Tomas Mustelin!, Lars G. Crabo?

| San Diego Natural History Museum, San Diego, California; 11904 Tallwood Court, Maryland 20854,
USA 2 Washington State University Adjunct Faculty; 724 14th Street, Bellingham, Washington 98225

Corresponding author: Zomas Mustelin (tmustelin@burnham.org)

Academic editor: D. Lafontaine | Received 14 March 2015 | Accepted 12 May 2015 | Published 15 October 2015
http://zo0bank.org/05826BC1-2746-4BAE-97EF-5BCO6BD63D5C

Citation: Mustelin T, Crabo LG (2015) Revision of the genus Aseptis McDunnough (Lepidoptera, Noctuidae, Noctuinae,
Xylenini) with a description of two new genera, Paraseptis and Viridiseptis. In: Schmidt BC, Lafontaine JD (Eds)
Contributions to the systematics of New World macro-moths VI. ZooKeys 527: 57-102. doi: 10.3897/zookeys.527.9575

Abstract

The genus Aseptis McDunnough (Lepidoptera, Noctuidae, Noctuinae, Xylenini, Xylenina) is revised to in-
clude 15 species based on morphological and molecular data. Several new synonymies are introduced. In
addition, two genera are described because of significant morphological differences from Aseptis: Paraseptis
gen. n., and Viridiseptis gen. n., resulting in the new combinations Paraseptis adnixa (Grote), comb. n.,
and Viridiseptis marina (Grote), comb. n. Although this work is primarily based on morphological data,
DNA sequence data for the 658-base pair “barcode” segment of the mitochondrial gene for subunit 1 of
cytochrome c oxidase was used as a secondary support for taxonomic changes within Asepzis and for the

two new genera. Our work should provide clarity and stability in a previously difficult genus.

Keywords
DNA barcode

Introduction

The genus Aseptis was described by McDunnough in 1937. A typical feature of the
genus is the indentation of the outer margin of the hindwing beneath its apex between
veins M1 and M3. Most species are rather dull gray or brown, many with diffuse mac-

Copyright Tomas Mustelin, Lars G. Crabo. This is an open access article distributed under the terms of the Creative Commons Attribution License
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58 Tomas Mustelin & Lars G. Crabo / ZooKeys 527: 57-102 (2015)

ulation in darker brown or black. Several species display marked variation between dif-
ferent geographical areas, as well as within any given locality. This often correlates with
the habitat and tends to yield paler and more diffusely marked individuals in dry and
sandy habitats, and darker more contrasting specimens in moist and lush habitats such
as in the Pacific Northwest. Many of these forms were described as separate species,
resulting in many more names than true species. From a total of 31 published species
names the latest checklist of North American Noctuoidea (Lafontaine and Schmidt
2010) contains 24 species, including “Aseptis” marina (Grote), which is associated
tentatively with Asepztis. In this revision the number of species is reduced further to 17,
of which15 are retained in Aseptis and two are placed in new genera.

Most species of Aseptis were described in the late 1800s and the first two decades of
the 1900s, with exception of four recent ones from southern California (Mustelin et al.
2000, Mustelin 2006). At the time of these older descriptions, the West was a frontier
and entomological collecting was restricted to a few localities such as Pacific ports, Provo,
Utah, a few localities in Colorado, and scattered sites sampled during geological expedi-
tions. Hence it is not surprising that short series from disparate sites were described as new
species without more thorough comparisons or anatomical examination. Indeed, Aseptis
binotata (Walker) was described seven times including in three different genera in a single
publication in 1865. The material accumulated in public collections over the last 100 years
now reveals that many species are polymorphic, while others are remarkably constant.

The combination of too many names, geographical variation, and the fact that many
Aseptis are similar gray-brown moths with diffuse markings has given rise to consider-
able confusion in public collections. A lack of published illustrations of most species
since Barnes & McDunnough’s publications a century ago (Barnes and McDunnough
1912a) has also contributed to a lack of clarity. In this revision, we define the status
of the species of Aseptis, describe two new genera for species previously associated with
the genus, and illustrate one or more representative adult specimens and the genitalia

of all of them.

Materials and methods

Wing pattern and genitalia structure terminology follow Lafontaine (2004). Terms not
defined in this reference are the penicillus, a broad-based lateral extension of the tegu-
men near the attachment to the vinculum (Forbes 1954), and the term “postreniform
patch” which we introduce for a pale area abutting the lateral reniform spot in the
distal medial and adjacent postmedial areas of the forewing (Fig. 1).

The male and female genitalia were prepared using standard methods (Hardwick
1950, Lafontaine 2004). Briefly, the detached abdomen was soaked in 10% KOH to
dissolve soft tissues. Dissection was performed initially in water followed by hardening
with isopropyl alcohol or ethanol. The male vesica and female bursa were inflated.
The preparations were stained with Chlorazole Black or orcein and were mounted in
Euparal on glass slides.

Revision of the genus Aseptis McDunnough....

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Tomas Mustelin & Lars G. Crabo / ZooKeys 527: 57-102 (2015)

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Revision of the genus Aseptis McDunnough.... 61

postreniform patch

Figures 1-2. Characters and nomenclature of the genus Aseptis. 1 Forewing of Aseptis binotata 2 hind-
wing of male Aseptis fumosa. The arrow points at the indentation typical of Asepztis.

The 658 base pair DNA “barcode” region of the mitochondrial cytochrome c oxi-
dase subunit 1 (CO1) was used to assess molecular variation of the species included in
Aseptis and related genera in recent check lists. Previously submitted samples available
as of February, 2015 at the Barcodes of Life Campaign (BOLD) at the University of
Guelph (Ontario, Canada) had been analyzed by standard DNA extraction, amplifica-
tion, and sequencing protocols for the BOLD initiative as described by Hebert et al.
(2003). The barcode sequences were compared using similarity trees obtained using
the Kimura-2-Parameter distance model as implemented on the Barcode of Life Data
Systems website (http://www.barcodinglife.org). Table 1 lists the major haplotypes of
specimens examined in this study.

This study is based on examination of material, including type specimens and
genitalia preparations, in the following collections:

AMNH The American Museum of Natural History, New York, New York, USA

BMNH _ The Natural History Museum [British Museum of Natural History],
London, UK

CNC Canadian National Collection of Insects, Arachnids, and Nematodes,
Ottawa, Ontario, Canada

FMNH Field Museum of Natural History, Chicago, Illinois, USA

LACM Los Angeles County Museum, Los Angeles, California, USA

LGC Lars Crabo Collection, Bellingham, Washington, USA

MSU Michigan State University, East Lansing, Michigan, USA

ODAC = Oregon Department of Agriculture, Salem, Oregon, USA

OSAC Oregon State Arthropod Collection, Corvallis, Oregon, USA

RHLC Ronald H. Leuschner Collection, now at the McGuire Center, Gainesville,
Florida, USA

SDNHM_ San Diego Natural History Museum, San Diego, California, USA

TEDC Thomas E. Dimock Collection, Ventura, California, USA

TMC
UCR
USNM
WFBM
WSUC

ZMH

Results

Tomas Mustelin & Lars G. Crabo / ZooKeys 527: 57-102 (2015)

Tomas Mustelin Collection, Potomac, Maryland, USA

University of California at Riverside, Riverside, California, USA
National Museum of Natural History [formerly United States National
Museum], Washington, District of Columbia, USA

W. F. Barr Entomological Collection, University of Idaho, Moscow, Idaho,
USA

James Entomological Collection, Washington State University, Pullman,
Washington, USA

Zoological Museum, Helsinki, Finland

Key to genera included in Aseptis McDunnough sensu Lafontaine & Schmidt, 2010

Male vesica with apical long spine-like cornutus; posterior half of female duc-
tus bineSae im erm braOUs anos cy Heecspseieeee tees ener teers Aseptis
Male vesica apex lacking cornutus or with minute cornutus; posterior half of
female ductus: bursae-at least-patrially sclerotized... ccz..cnnsecesesenesetceneeesbetes 2
Ampulla of male clasper present; female corpus bursae with four long signa
and posterior ductus bursae sclerotized circumferentially ............. Paraseptis
Ampulla of male clasper absent; female corpus bursae lacking signa and pos-
terior ductus bursae with sclerotized plate in ventral wall........... Viridiseptis

Key to the genus Aseptis McDunnough

TA let reese ath caeh tena nen teak ys Seonan Poms tensae Mee erh beaseet¥ Acne Raden ty lsn Case Pilko ln Boose 2
PETIA x Ais terest te ach tects sock techie teat A deal el tee LIE than ttl ttl tt hal abit dean 19
Pans UUDE LSA RET Kote eee RIERA RAED) De ace A AOD os Ted DO A. serrula
gee Wal URES MILM ROTH dh ROM Oech Sa Ee SN SR EN, SPE ro arte Semen a Fd 3
TPIS AD SIE (5. Me na ate tra teks nano banutBohinecnishes Ut shea tn banites Sete teat sbellratehl weslubad 4
IDISILUS elOM Gate, MATLOW ecu ceconer wecyeusecereeneecyesnecaneevonncadvrvecanvenecetnivenssicensels 8

Ventral cucullus with spike-like process; forewing mottled olive green to olive
yellows, Clalit G thilair csr. cask emtnas ros ieee aT ATR GNEAE 5
Ventral cucullus normal, rounded; forewing not olive; widespread, including
(RENIN COTM ee Toei ieotins os Ahmar lene. A ae so hye AR oO lens a et ae 6
Forewing ground color olive green with yellow tan and black pattern; Kern
and Tuolumne counties, California, and north; male valve nearly straight;
female genitalia indistinguishable from A. pseudolichena ..........104+. A. lichena

Revision of the genus Aseptis McDunnough.... 63

Forewing ground color light olive to olive-yellow; Kern and Tuolumne coun-
ties, California and south; male valve bent slightly ventrad at mid-point; fe-
male genitalia indistinguishable from A. lichena...........1- A. pseudolichena
Vesica with single cornutus; forewing mottled light yellow tan and gray; de-
sents of southern Califoriia atid ATIZONAA,.<.c.ssv01-0¥000¢enesecesseee ences A. catalina
Vesica with two or more cornuti; forewing not as above; widespread in west-
SAU NORCM TCT Ca 4 NT... ocean Mea ee a eee oa Ee Ary ae ee A De Bl a 7
Valve slightly S-shaped; forewing mottled gray, or gray and tan...A. characta
Valve nearly straight; forewing dark, blackish oo... eee A, fumosa
Digitus perpendicular to valve; southern California... cece eeeeeeeeeeeee 9
Digitus oblique to valve, pointed ~45° toward ventral cucullus; widespread,
includine southern Cali Omnia ki coiva iene baucteon else wdesathetommavioninliwanebzerne: 10
Digitus origin near ventral valve with most of it below ventral valve margin;
forewing dark brown to black brown; widespread in southern California......
sfdw ie sel pe wea lveet ralet i Uanwcdl ye ewiiWe rites AEs we cnedy MRAM tutto tnhdap A. perfumosa
Digitus origin on mid-valve near base of ampulla, barely reaching ventral
margin; immediate coast near San Diego, California ................ A, torreyana
Aedeagus longer, > 4x as wide as long; smaller narrower-winged species
(wingspan < 35 mm); forewing with contrasting light postreniform patch or
sinalliblacebasalncas in eye eet ocr Sees eee eee een ee 11
Aedeagus stout, < 4x as wide as long; large broad-winged species (wingspan
> 35 mm); forewing without basal dash and usually without postreniform
VOY 181 # MECN, era A ew ele. Uae at Sr. ope Aseptis fumeola species group...12
Small basal dash present, evident in all but the darkest specimens; forewing a
shade of brown; hindwing base gray; widespread in western North America...
Re Ape REN, GC RSS sate od AEN eg sa eRe RE Bo ean Bods WOR, ee le ere A. binotata
Basal dash absent; forewing gray with patches of pale rusty brown; hindwing
base white with streaks extending distally; deserts of Southwest and southern
© alienated. tse or one as Pena maa ee as ene caahec. hoa earl e A, susquesa
Forewing smooth gray with pale costa; maculation reduced to dark filling of
spots and dotted lines; extreme southern California... A, murina
Forewing not as above, costa similar to rest of wing; West Coast and parts of
Southwest, including southern-Galiformia... c.cdeedseossencdecbuceonnstdocnassstsonadtale 13
Forewing mottled gray brown with conspicuous gray filling of spots, small
yellowish postreniform spot, and irregular black to dark gray shade proximal
to subterminal line; California, Arizona, and southern Utah and Nevada.....
ee Sib ai aaa ar aacsenag aan ener OR eRe tives fod nian coven ene onns del ag tag rates A. fumeola
Forewing light or dark but more uniform, filling of spots not strongly con-
trasting, medial area between reniform spot and postmedial line not signif-
cantly lighter than rest of wing or reddish in central and northern California;
Galitoriia, Arizona, Oreson, arid: Washi ne toni. ..tssccecrmrevevtevsevoaaernestvence 14
Forewing ground: color strongly red DOWN ..sisse.cscsscsseasseoctstsnstececestevessuees 15

Tomas Mustelin & Lars G. Crabo / ZooKeys 527: 57-102 (2015)

Forewing not red brown, sometimes patchy reddish areas near reniform spot

in-centtaland northern, Califonniants..225.:.ibs.reltesselectueck ince sevsbaeetelgseleed 18
Forewing rusty red brown with darker markings; extreme southern California ..
brea sses ora tcuanasdiat. ecaccecs bee EIEEE Dhocassbook treme mR cesar on ee A, ferruginea
Forewing bright red brown, occasionally with darker markings; San Benito
County; Calitorintae om sores oo tee sonar eee tee A. fanatica (part)
SOC SEN alta 8 ca. eee de Sus OSs dee wn ocean eens aoe eee ety 17
Ceiitral-CalitOritiastO gy ASHI CLO: woes l ake, wseuttesl Oeics osc bal Mend Siete! 18

Forewing ground color slightly mottled pale gray brown; male valves oriented
> 120° relative to each when mounted flat; female corpus bursae ~5x as long
BS GAEL a usuoh's Nv belen cise Oe Sai Ne Sonat Bs Oe du, cots, Bee Ah A. ethnica (part)
Forewing ground color medium to dark brown; male valves oriented at ~90°
when displayed similarly; female corpus bursae ~7x as long as wide..............
eg at EB Pes 5 Oe toe cree Ath clo ld hat Ah aie eR cc A, fanatica (part)
Forewing ground color slightly mottled brown, usually with evident dark
shade preceding subterminal line and often with reddish postreniform spot;
male valves oriented = 120° relative to each when mounted flat; female corpus
Duster s5 x as TOM eas Wie, 4 Sch stech delsciteuatellay aghcuenuahobalaaets A. ethnica (part)
Forewing ground color blackish brown, maculation faint; male valves ori-
ented at ~90° when displayed similarly; female corpus bursae ~7x as long as
SUES Ca {cells AP a a 7h refrac re te Sana RR aie one ent Ae ern A. fanatica (part)
Corpus bursae elongate, 5—7x as long as wide, with small cone-shaped ap-
pendix bursae; papilla analis without long hair-like basal seta... eee
gt hag Be 2k Ne se PR Le Nee See Aseptis fumeola species group...12
Corpus bursae wider, ovoid, 1.3—1.5x as long as wide, with appendix bursae

not as above; papilla analis with sparse or thick hair-like basal setae.......... 20
Forewing mottled olive green to yellow green; California... eee 5
Forewing ground color gray or brown; widespread, including California... 21
Apex of papilla analis with a thin sclerotized flange.................. A. perfumosa
Apex of papilla analistlackingca Mange .. tlt wos Pusocwsbeatavas caer sbormsans eshopetaa. 22,

Hairs at base of papilla analis dense, with expanded ventral patches; South-
west-deserts:and southern Californias on Ws.cz.n caeeess cubhs Beeson dense te vae estates 23
Papilla analis with a sparse single row of basal hair-like setae; widespread,
iitcleine-southerng@aliornia ih: F.we... 3. truer snnstaracdlna tase eerenghaeaene nae taak 24
Papilla analis covered with short needle-like setae, medial dorsal margin
smooth; forewing patchy light yellow tan and gray ...... eee A, catalina
Papilla analis rugose, scale-like, dorsal medial margin irregular; forewing gray
SAT 0 Cesta Fea e Os vole 1d cccupteory an tape ere Irie RRA RRA RT A, serrula
Ground color of forewing dark smoky brown to nearly black; appendix bur-
sae bluntly rounded and mediolaterally compressed...........::.::0000+ A. fumosa
Ground color variable, brown to gray; if dark brown then more light scaling
including filling of lines and ochre postreniform patch; appendix bursae not
ASADOVR PAST MAM C UIC, PE REN Gestalt le. uun dane tcustt ssnarte oi eel neaee as 25

Revision of the genus Aseptis McDunnough.... 65

25 Ground color of forewing a shade of brown (including gray brown), but lack-

I OPERIENS IV CoC ta YAP CAST otha sina csbaattl atu andee natal ua aeatrsd ctu A. binotata
- Forewing gray or gray with light tan or rusty accent ...... cs eeeseseeeteereeeees 26
26 Forewing mottled light gray, with at most minor patches of olive or tan

Sar | (cx Pea AREA ETOORAER AR dae PRUE MIR, Ae Ree RAD aoe eT A Te A RRA TTS SS 27,
- Forewing gray with extensive tan or rusty-tan scales... eeeseseceseereeeeee 28
27 Forewing mottled light gray, markings include a thin black basal dash; Pacific

Coast near San Diego, California; female genitalia unknown....A. torreyana
- Forewing mottled light and medium dark gray to darker medium gray, black
dash absent; widespread, including in southern California; appendix bursae
broad based but short; corpus bursae lacking signa .......... A. characta (part)
28 Forewing pattern longitudinally streaked, with rusty patches that are most
prominent in fold and distal to reniform spot; bursa copulatrix with corpus
bursae and appendix bursae nearly equal in size; desert Southwest and south-
SUT (Pali eM at ae Beet Oe ese Oo Se Dine sal ey Pou aot Aa age Sane Asad A. susquesa
- Forewing pattern mottled gray and tan or rusty tan, not streaky; appendix bur-
sae much smaller than corpus bursae; widespread in western North America ...
PRRURTINT oe bt svevTre To EE eveN on, Lane IT 4/7 FT REL Se A. characta (part)

Aseptis McDunnough, 1937

Aseptis McDunnough, 1937: 59.

Type species. Hadena genitrix Grote (a synonym of Aseptis binotata (Walker)) by origi-
nal designation.

Diagnosis. Aseptis is a moderate-sized genus of medium-sized noctuids (wing-
span 27.5—45.0 mm) in the subtribe Xylenina Guenée of the tribe Xylenini Guenée
of the subfamily Noctuinae Latreille (Lafontaine and Schmidt 2010) from western
North America. Adults are typically dull mottled gray or brown, although a few spe-
cies are red brown or nearly black, with typical noctuid lines and spots, often with
a pale patch in the medial and postmedial areas abutting the lateral reniform spot
(“postreniform patch” (Fig. 1)), which is easily mistaken for the reniform spot. The
reniform, orbicular, and claviform spots are present in most species and are often large
and closely positioned; the reniform spot is usually figure-eight shaped. The hindwing
outer margin is concave between M1 and M3, M2 is visible and is closer to M3 than
M1, and the wing is often palest with loss of scales between M1 and M3 (Fig. 2).
The male antenna is filiform, serrate in Aseptis serrula (Barnes & McDunnough). The
male abdomen has basal coremata with pockets extending on segments one and two.
The male genitalia have a narrow sharply-pointed uncus; a tegumen that is laterally
compressed near the uncus base and has broad penicillus lobes; a strap-like valve
with small sacculus and weakly differentiated rounded cucullus with a weak corona
(ventral cucullus pointed in the Asepztis lichena species group), a curved ampulla of the

66 Tomas Mustelin & Lars G. Crabo / ZooKeys 527: 57-102 (2015)

clasper oriented perpendicular to or parallel to the costa, and an elongate triangular
or spike-like digitus arising from a weakly sclerotized plate on the mid-valve (digitus
absent in several species); the aedeagus is tubular with slight ventrad bend distally,
and the vesica is 114-2 x aedeagus length with 90—180° bend ventrad at the base, and
bears a long proximally-directed apical cornutus and additional 0—2 smaller cornuti
and 2—3 broad-based diverticula. In the female, the papilla analis is lightly sclerotized,
triangular, with a rounded tip, and is covered with short spike-like setae (rugose scales
in A. serrula) and from zero to innumerable hair-like basal setae; the ductus bursae
is membranous except near the corpus bursae; the moderately-sclerotized appendix
bursae is sack-like or weakly bilobed and extends posteriorly from the left ventral
corpus bursae; the corpus bursae is ovoid, 1.3—7 x as long as wide, with 0—4 long
narrow signa.

Aseptis can be distinguished from all genera other than Paraseptis Mustelin & Cra-
bo and Viridiseptis Mustelin & Crabo, both described below, by the concave hindwing
notch. Aseptis males have long apical cornutus on the vesica, absent or very small in the
other genera. Aseptis females lack sclerotization of the posterior ductus bursae.

Distribution and biology. Aseptis species mainly occur west of the Great Plains
from south-central Alberta and southern British Columbia to northwestern Mexico;
one species, Aseptis characta, extends eastward into the Great Plains as far as Manitoba.
The greatest concentration of species is near the Pacific Coast, particularly in southern
California, and in the desert Southwest. The adult flight season is from late spring
to late summer and is often fairly long, but species in desert habitats usually fly only
during the spring. They occur in a variety of habitats from forest, shrub steppe and
chaparral, to desert. As typical of the tribe Xylenini (Fibiger and Lafontaine 2005), the
larvae feed on the leaves of woody plants.

Discussion. Fifteen Aseptis species are recognized herein. Seven sort easily into
species groups, five in the Aseptis fumeola species group and two in the Aseptis lichena
species group. No natural groupings were found for the other species.

Two species previously associated with Aseptis differ significantly in structure from
the other members of the genus. “Aseptis” marina was moved recently to Aseptis from
Oligia Hiibner in the Apameini (Lafontaine and Schmidt 2010). They noted that al-
though it appears be related to Aseptis they are not congeneric, adding the quotations
to denote the tentative association. Its hindwing margin has a slightly concave segment
like those of Aseptis, but the male and female genitalia are strikingly different. We
describe Viridiseptis for it below and introduce Viridiseptis marina (Grote), comb. n.
Its relationship to the Xylenini is also discussed further. The other outlier, Paraseptis
adnixa (Grote), comb. n., is surprising because it superficially resembles A. binotata
but differs from Aseptis in several features of the male and female genitalia. In addition
to the anatomic differences, the CO1 barcodes of V. marina and P. adnixa variably sort
away from Aseptis and each other within a large selection of Xylenini.

The name Aseptis was presumably chosen by James McDunnough to denote its
distinction from Septis Hiibner, a synonym of Apamea Ochsenheimer, in the Apam-
eini. Aseptis and Apamea are not related closely.

Revision of the genus Aseptis McDunnough.... 67

Aseptis fumeola species group

The Aseptis fumeola species group consists of five species (Aseptis fumeola (Hampson),
Aseptis ethnica (Smith), Aseptis murina Mustelin, Aseptis ferruginea Mustelin, and Asep-
tis fanatica Mustelin). Its members are relatively large (wingspan = 35 mm) with broad
forewings with diffuse markings. The hindwing is dark, which obscures the veins. The
male valve is long and narrow with a truncated or foot-shaped cucullus, upright am-
pulla, and oblique pointed digitus. The aedeagus is stubby, usually 3—4x as long as
wide, with a large sack-like vesica with large diverticula and long apical cornutus. Fe-
males have bursae with an elongate corpus bursae and small appendix bursae, and lack
basal hairs on the papillae anales. The genitalia are similar within the group and the
species are most easily identified by their habitus.

Aseptis ethnica and A. fanatica display geographical variation in that both are much
darker at the northern than at the southern ends of their ranges. Nevertheless, mem-
bers of this species group tend to be relatively constant in appearance at any given
location. An exception to this is a population of A. fanatica from San Benito County,
California, in which approximately half of the specimens are bright reddish whereas
the other half are of the typical black color.

All species in the Aseptis fumeola species group occur in California. The ranges of two
of the species extend north to the Pacific Northwest, two as far East as Arizona, and two
are restricted to southern California. All five species are sympatric in southern California.

The larvae of A. fumeola, A. ethnica, and A. fanatica feed on Manzanita (Arcto-
staphylos spp., Ericaceae) and related plants.

The CO1 barcodes of the species in this species group are relatively similar and
cluster tightly within the genus. The largest difference in barcode sequences between
two species in the group is 2.4%. Aseptis fanatica is the only species in the species group
with more than a single barcode haplotype.

Aseptis fumeola (Hampson, 1908)
Figs 3, 4, 59, 76

Trachea fumeola Hampson, 1908: 186.
Trachea (Hadena) probata Barnes & McDunnough, 1910: 153.

Type material. Trachea fumeola: Holotype male [BMNH, photograph examined].
Type Locality: Pinal Mountains, Arizona. Trachea (Hadena) probata: Holotype fe-
male [USNM, photograph examined]. Type Locality: Huachuca Mountains, Arizona.

Diagnosis. A large and broad-winged Aseptis, wingspan 41.1+1.6 mm (n=25; range
38.5-45.0 mm), with a dark slightly shiny gray-brown forewing with contrasting dark
patches, particularly in the basal and postmedial areas. The medial area typically is paler
with reddish tan near the conspicuous large black reniform spot, small round orbicular
spot, and short claviform spot. The reddish postreniform patch is relatively prominent

Tomas Mustelin & Lars G. Crabo / ZooKeys 527: 57-102 (2015)

‘(VD “0D omuag UPS) seuroj
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Revision of the genus Aseptis McDunnough.... 69

for the species group. The postmedial line usually is well marked and curves around the
reniform spot. The serrated subterminal line is a prominent border between the postme-
dial and paler subterminal areas. The hindwing is smoky gray brown, darker in females.

Aseptis fumeola can be identified by its large size and patchy forewing maculation.
Some northern California A. ethnica have a similar brown forewing with reddish tan
around the spots; A. fumeola tends to be more contrasting, less reddish, and a bit larger.
Occasional Aseptis perfumosa specimens are dark brown with reddish suffusion in the
upper medial area. This form is always much smaller and darker than A. fumeola, has
a narrower forewing, and much different genitalia. The male and female genitalia are
as in the description to the species group. The male valve has a foot-shaped cucullus.

Distribution and biology. Aseptis fumeola is known from Arizona, southern and
central California, southern Nevada, and south-eastern Utah. It flies in the foothills
and mountains in dry chaparral, parkland, and conifer forest. Most records are from
June and July. The larva and pupa were described and figured by Comstock (1940a).
The pale-green larvae were found and reared on new leaves of manzanitas (Arctostaphy-
los spp.). Crumb (1956) found it on A. pungens Kunth in Arizona and described the
larva as strange and sluggish.

Remarks. ‘The Latin name fumeola means smoked, likely to denote its black-pep-
pered maculation.

Aseptis ethnica (Smith, 1899)
Figs 5-8, 60, 77

Hadena ethnica Smith, 1899: 263.

Type material. Holotype male [USNM, examined]. Type Locality: Yosemite, Cali-
fornia.

Diagnosis. Aseptis ethnica is a large and broad-winged, wingspan 41.2+1.5 mm
(n=25; range 38.0-43.5 mm) with ill-defined markings. In southern California its
forewing is dull grayish tan with a grainy appearance (Figs 5 and 6). In central and
northern California and Oregon, it is dull deeper brown, sometimes with some red-
dish tones surrounding the dark-filled reniform and orbicular spots (Figs 7 and 8). The
holotype from Yosemite is of the darker form and is reminiscent of A. fumeola. Most
A. ethnica are much less contrasting than A. fumeola and lack its contrasting black-
outlined spots, patchy dark shading, and reddish postreniform patch. Aseptis ethnica
can be challenging to distinguish from A. fanatica, particularly when worn. Typically,
A. ethnica is larger with a broader forewing, has a grainier forewing pattern, and is the
lighter species in any location. In southern California, A. ethnica is pale grayish tan
whereas A. fanatica is darker gray brown. In northern California, A. ethnica is as dark
as A. fanatica in San Diego County, but in this area A. fanatica is nearly black. In San
Diego County, A. ethnica can be distinguished from A. murina by being grainier and
browner, and by lacking its characteristic pale costa.

70 Tomas Mustelin & Lars G. Crabo / ZooKeys 527: 57-102 (2015)

The male genitalia of A. ethnica can be distinguished from those of A. fanatica by
the angle subtended by the two valves as noted in the key. The female bursa is slightly
shorter in A. ethnica than in A. fanatica, approximately 5x its width in A. ethnica and
7x in the latter species.

The CO1 barcode DNA sequence of A. ethnica is closest to A. murina, differing
by 1.8%.

Distribution and biology. Aseptis ethnica is known from Arizona, California,
western Oregon, and Baja California Norte, Mexico. The northern limit is not known
precisely due to similarity of this species and A. fanatica in this portion of its range;
however, it occurs at least to Jefferson County, Oregon, based on DNA. Aseptis eth-
nica flies in open Pine and Oak forest and mountain chaparral, mostly at elevations of
above 1500 m in southern California but at lower elevations farther north. It has been
found from early May to August and can be locally abundant during its peak flight in
June and July. In the mountains of southern California (e.g., Laguna, Volcan, Palo-
mar, and Santa Rosa Mountains) it often flies together with A. fumeola, A. fanatica,
and occasional specimens of A. ferruginea. The immature stages are unknown, but the
larval food plant was revealed by J.B. Smith’s description of the species in 1899 based
on adult specimens raised from caterpillar on manzanitas (Arctostaphylos spp.).

Discussion. The name et/nica is Latin and means heathen. Perhaps Smith was
inspired by the raw wilderness of Yosemite in late 1899.

Aseptis murina Mustelin, 2000
Figs 9, 10, 61, 78

Aseptis murina Mustelin, 2000. In: Mustelin et al. 2000: 8.

Type material. Holotype male [SDNHM, examined]. Type locality: Inaja Picnic
Ground, San Diego County, California.

Diagnosis. ‘This is a large species with perhaps the broadest forewing in the group,
wingspan 40.0+1.0 mm (n=12; range 39-42 mm). The forewing is smooth gray with
a slightly bluish sheen when fresh, a diffuse and faint dark reniform spot, postmedial
line of black dots, and a pale-cream costa in fresh specimens. The hindwing is barely a
shade paler than the forewing.

This species can be identified by the combination of large size, smooth gray fore-
wing with pale costa, and southern California distribution. It is most likely to be con-
fused with A. ethnica, especially when worn.

Distribution and biology. Aseptis murina is known only from southern California
where it occurs in coastal chaparral, foothills, mountain brush land and oak forest, and
in the mountain-desert transition zone from sea level to 2000 m. It can be found with
all other species of the species group. Localities for it include the Cuyamaca, Laguna,
and San Gabriel Mountains, Pinyon Crest, and San Marcus Pass in Santa Barbara
County. The flight period is from early May to July. The early stages are unknown.

Revision of the genus Aseptis McDunnough.... 71

Discussion. The name murina is Latin and means mouse-like to denote the
smooth murine appearance of the moth.

Aseptis ferruginea Mustelin, 2000
Figs 11,12;-62, 79

Aseptis ferruginea Mustelin, 2000. In: Mustelin et al. 2000: 8.

Type material. Holotype male [SDNHM, examined]. Type locality: Wynola, San
Diego County, California.

Diagnosis. Aseptis ferruginea is one of the smallest and most narrow-winged spe-
cies in the group, wingspan 36.4+0.5 mm (n=6; range 35.5-37.0 mm). Its brown fore-
wing is distinctly reddish, deep claret when fresh and rustier when worn, and the veins
are usually black. Most specimens have a clearly-marked dark-filled reniform spot and
a jagged pale subterminal line. The hindwing is distinctly paler than the forewing. The
male valve differs from others in the species group in being even and straight with a
perpendicular lateral cucullus that is straight or slightly concave.

Aseptis ferruginea can usually be identified by superficial appearance, especially its
red color, and males can be confirmed by dissection. Some central California A. fa-
natica are bright red brown and could be confused with it, although this morph is not
known from within its geographical range; these A. fanatica are smoother than A. fer-
ruginea and lack the other forewing markings described above.

The CO1 barcode sequence of A. ferruginea is the most unique of any member
of the species group. It is closest to that of A. ethnica from which it differs by at
least 2.2%.

Distribution and biology. This species is endemic to southern California. All
records are from San Diego County from an area between Boulevard-Manzanita near
the Mexican border north to Lake Henshaw at altitudes of 800-1600 m. It flies in
open oak forest, foothill chaparral, and in the mountain-desert transition zone. Aseptis
ferruginea can be encountered together with the much more abundant A. ethnica and
A, fanatica at higher altitudes and with A. murina at lower elevations. Records are from
late June to August. ‘The early stages are unknown.

Discussion. The name ferruginea is Latin and means rusty.

Aseptis fanatica Mustelin, 2006
Figs 13-18, 63, 80

Aseptis fanatica Mustelin, 2006: 27.

Type material. Holotype male [SDNHM, examined]. Type locality: Pine Cove, San

Jacinto Mountains, Riverside County, California.

72 Tomas Mustelin & Lars G. Crabo / ZooKeys 527: 57-102 (2015)

Diagnosis. This species is similar in size and shape to A. ferruginea, wingspan
38.9+1.6 mm (n = 25; range 35—42 mm). In southern California A. fanatica has a dark
chocolate-brown forewing (Figs 13 and 14), whereas in northern California, Oregon,
and Washington it is darker brown to nearly black (Figs 15 and 16). Some individuals
in central California are smooth bright red brown (Figs 17 and 18). The maculation
is dark, either diffuse or weakly contrasting. The most prominent markings are the
black-filled reniform spot, a black shade proximal to the incomplete pale subterminal
line, and pale-yellowish spots on the costa at the antemedial and postmedial lines.
Well-marked specimens have a serrate black postmedial line, some black on the veins,
and scattered pale scales giving them a peppered look. The hindwing is slightly paler
than the forewing, particularly in males.

Separating A. fanatica from A. ethnica can be challenging. As a rule, A. fanatica
is the darker species at any location. In southern California A. fanatica is dark gray
brown whereas A. ethnica is pale tan gray. In northern California where A. ethnica is
darker and often has some reddish brown around the spots, A. fanatica is nearly black.
Aseptis fanatica tends to be narrower winged and smaller, but there is overlap in size. If
necessary, the genital characters of the male valves and female bursae given in the key
to species can be used to distinguish the two species.

Aseptis fanatica is the only species in the species group with two CO1 barcode hap-
lotypes, these separated by 1.3%. Specimens with both haplotypes are found through-
out its range and display no consistent differences in habitus or male or female geni-
talia. Similarly, the distinctive red morph from San Benito County flies with typical
black specimens with which they are indistinguishable by barcodes or genitalia.

Distribution and biology. Aseptis fanatica is known from Washington, Oregon,
California, and Baja California Norte, Mexico. It flies in many different habitats like
brush land and open forest in southern California mostly at 1000—2000 m but occurs
at lower elevations farther north. The flight period is from early May to August in the
south and in mid-summer in the Cascades. It can be abundant. In the Pacific North-
west, A. fanatica feeds on species of Ericaceae such as madrone (Arbutus menziesii
Pursh.) and various species of manzanitas and bearberry (Arctostaphylos spp.) (Miller
& Hammond 2003, as A. ethnica). Bearberry (Arctostaphylos uva-ursi (L.) Spreng.) is
probably the only suitable foodplant for this species in the Washington Cascades.

Discussion. Prior to its description in 2006, this species was thought to represent
the southern California form of A. ethnica. In fact, the holotype of Hadena ethnica is
quite similar to A. fanatica from San Diego County.

The mixed red and black population from San Benito County is a unique phe-
nomenon. The red color might be due to a gene mutation, but could also be adaptive.
Many noctuids that feed on Arctostaphylos as larvae are a similar red color, including
Mesogona rubra Hammond & Crabo in the subtribe Xylenina.

The name fanatica means fanatic and was selected as the antithesis of ethnica (hea-
then). At the time, fanatic had a less sinister meaning than it does in today’s world.

Revision of the genus Aseptis McDunnough.... 73

Aseptis lichena species group

The Asepztis lichena species group consists of two closely related species from the moun-
tains of central and southern California. They are unique in the genus in that the fore-
wing is mottled olive, darker olive green in Aseptis lichena (Barnes & McDunnough)
and paler yellow tan in A. pseudolichena Mustelin & Leuschner. The male valves have
a unique acute spine from the ventral cucullus and lack a digitus. The vesicas have a
single long apical cornutus. The female bursae are indistinguishable, with large corpora
bursae without distinct signa and rounded appendices bursae. The CO1 barcode se-
quences of the two species differ by approximately 2.3%.

Aseptis lichena (Barnes & McDunnough, 1912)
Figs 19, 20, 64, 81

Andropolia lichena Barnes & McDunnough, 1912b: 17.

Type material. Holotype female [USNM, examined]. Type locality: Deer Park
Springs, Lake Tahoe, California.

Diagnosis. A medium-sized noctuid with a wingspan of 35.5+1.8 mm (n=8; range
33-39 mm) with a powdery dark olive-green forewing produced by a mixture of black,
green, and yellow scales. Aseptis lichena is darker green than A. pseudolichena. The male
valve of A. lichena is nearly straight whereas that of A. pseudolichena is bent slightly
ventrad at its midpoint. In practice, most specimens can be assigned to a species based
on geography, except in an area of overlap at the south end of the Sierra Nevada in
Kern and Tuolumne counties. Aseptis lichena occurs to the north of this zone. Females
are best associated with the males.

Distribution and biology. ‘The relatively few specimens of A. lichena we have
examined are from south-central California (Tehachapi Mountain Peak, Kern Co.)
and north-central California (near Blairsden, Plumas Co., Lake Tahoe, and Yosemite
Park). It is also reported from Mount Shasta, Mount Lassen, and other locations in
northern California. It flies during mid-summer. ‘The early stages are unknown.

Discussion. This species was described in the genus Andropolia Grote, pos-
sibly because the holotype is a dark female with a relatively inconspicuous inden-
tation of the hindwing. Specimen labels found by the senior author suggest that
McDunnough suspected that this was incorrect. A female collected in Yosemite
National Park, Camp 19, on 15 July 1937 by F.L. Cramer has a second label:
“McD needs,” a third label: “Andropolia lichena B & McD., Det. Dr. J. McDun-
nough,” and a fourth label: “Probably misplaced in “Andropolia” — McD.” None-
theless, it remained in Andropolia until it was associated with Aseptis by Mustelin
et al. (2000).

Tomas Mustelin & Lars G. Crabo / ZooKeys 527: 57-102 (2015)

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qewsay ‘vuagoyopnasd sudasy y@ (WD “OD O8a1q] URS) seu ‘yuagoyopnasd sudasy EJ (YD “OD O8aIq ues) adAesreg aeuray ‘yuagotopnasd sudasy TZ (YD “OD S73
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Revision of the genus Aseptis McDunnough....

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76 Tomas Mustelin & Lars G. Crabo / ZooKeys 527: 57-102 (2015)

Aseptis pseudolichena Mustelin & Leuschner, 2000
Figs 21-24, 56, 82

Aseptis pseudolichena Mustelin & Leuschner, 2000. In Mustelin et al. 2000: 10.

Type material. Holotype male [LACM, examined]. Type locality: East Fork of Wood-
wardia Camp, San Gabriel Mountains, Los Angeles County, California.

Diagnosis. Aseptis pseudolichena is similar in size or slightly smaller than A. lichena,
wingspan 33.84+1.4 mm (n=25; range 30.5—36.0 mm) and resembles it closely. Aseptis
pseudolichena is more yellowish as a result of a mixture of pale tan or yellowish scales
and scattered tan, olive, and black scales. Some specimens are very pale yellow, others
pale tan, and some olive tan. The antemedial and postmedial lines are serrate, and the
postmedial line is followed by white and black dots on the veins. A pale subterminal
line is usually visible. The orbicular spot is round and filled with ground color, whereas
the reniform spot is large, outlined in black and filled with dark scales. A faint pale
postreniform patch is present. Males have a pale streak between hindwing veins M1
and M3.

Most A. pseudolichena can be separated readily from A. lichena by their pale-yellow-
ish to olive-tan color, which is darker olive tan in A. lichena; however, dark specimens
of A. pseudolichena may not be distinguishable without dissection. The male genitalia
are similar to those of A. lichena, but differ in that the valves are angled ventrad at mid-
length and the cucullus is smaller. Most specimens can be assigned to a species based
on locality as described under A. lichena.

Distribution and biology. Aseptis pseudolichena is endemic to southern California
with records from San Diego, Riverside, Los Angeles, Ventura, San Bernardino, and
Tuolumne counties. It may overlap with A. lichena in Kern and Tuolumne counties.
Aseptis pseudolichena is found in open pine and oak forest, open areas with grass and
scrub, and foothill chaparral. It seems to be most common on the desert side of the
mountain peaks, and can be locally abundant. It flies from June to August depending
on elevation. A number of specimens in the Los Angeles County Museum were raised
from larva on Ribes malvaceum Sm. (Grossulariaceae). The pupa was described and

figured by Comstock (1955) under the name Andropolia lichena.

Aseptis binotata (Walker, 1865)
Figs 25-34, 66, 83

Mamestra binotata Walker, 1865a: 663.

Miana rubiginosa Walker, 1865a: 675.

Hadena extersa Walker, 1865b: 728.
Taeniocampa paviae Strecker, 1874: 94, syn. n.
Hadena curvata Grote, 1874b: 157, syn. n.
Hadena genitrix Grote, 1878: 237, syn. n.

Revision of the genus Aseptis McDunnough.... Td.

Hadena inconspicua Smith, 1893: 142, nomen nudum
Hadena dilara Strecker, 1898: 7, syn. n.

Hadena bultata Smith, 1906: 228, syn. n.

Trachea cara Barnes & McDunnough, 1912c: 52, syn. n.

Type material. Mamestra binotata: Holotype male [BMNH, photograph examined].
Type locality: Vancouver Island, British Columbia. Miana rubiginosa: Holotype male
[BMNH, not examined]. Type locality: Vancouver Island, British Columbia. Hadena
extersa: Holotype male [BMNH, photograph examined]. Type locality: Vancouver Is-
land, British Columbia. 7aeniocampa paviae: Syntypes [Strecker coll., not examined].
Type locality: California. Hadena curvata: Holotype female [BMNH, photograph
examined]. Type locality: Mendocino, California. Hadena genitrix: Holotype female
[BMNH, photograph examined]. Type locality: Nevada. Hadena inconspicua: Lecto-
type male designated by Todd (1982) [USNM, examined]. Type locality: California.
Hadena dilara: Holotype female [FMNH, photograph examined]. Type locality: Col-
orado. Hadena bultata: Lectotype male designated by Todd (1982) [AMNH, exam-
ined]. Type locality: Glenwood Springs, Colorado. Trachea cara: Syntypes [USNM,
examined]. Type locality: Eureka and Provo, Utah.

Diagnosis. Aseptis binotata is a common medium-sized member of the genus with
a wingspan of 32.541.3 mm (n=25; range 29.5—35.0 mm). It is the most variable
Aseptis with respect to forewing color and pattern strength. It may be brownish, warm
dark brown, pale to medium gray brown, yellowish light brown, or reddish brown
depending on locality. The most noticeable marking in dark specimens is the large
pale yellowish postreniform patch, which is bisected by the dark postmedial line. The
antemedial line is strongly convex laterally and is filled with light tan. Black forewing
markings include a series of wedges near the outer margin below the apex, the outlines
of the three forewing spots, and in most specimens a short black basal dash. Pale speci-
mens can be washed out or have contrasting dark markings.

There is striking variation in this species, both within populations and over larger
distances. Specimens from the coastal region of the Pacific Northwest are fairly uni-
form with a warm dark brown forewing with distinct maculation (Fig. 25). Southern
California specimens of A. binotata range from nearly as dark (Fig. 26) as Northwest-
ern ones to pale reddish, tan, or pale gray brown with a less contrasting postreniform
patch (e.g., Fig. 27). These pale forms were described as Hadena curvata Grote and
Taeniocampa paviae Strecker. Populations from drier habitats east of the coastal moun-
tains also tend to be pale and more uniform in color, often with gray tones (Figs 28,
29). Populations from near the Rocky Mountains are also variable, mostly dull gray
brown (Figs 30, 31, but those from areas of Colorado, Utah, and New Mexico with
reddish substrate are yellow tan to orange tan, often with reduced dark patterns (Figs
32-34). These colorful morphs were described as Hadena dilara Strecker and Hadena
bultata Smith.

The male genitalia of A. binotata have a valve with a curved upright ampulla, a long
downwardly curving digitus, and a weak constriction at the base of the cucullus. The

Tomas Mustelin & Lars G. Crabo / ZooKeys 527: 57-102 (2015)

(YD “0D o8sIq ues) sfeur ‘vuLvUL sudasIp1s1A Bg (YD “OD O82Iq UeS) seu ‘vuLvUL sydastp1s1A Lg (Dg ‘A2;sueT) syeuT
‘vxIUpp sUdasBLDT 9g (WD “OD OAUT) 2JeWIDZ ‘vxzupv sudasBLUg GG (YD “OD 0821 UeS) seu ‘wx1Upy sudasvLD HG (YD “OD O82Iq UeS) IZewW ‘wxiUupy sudasvLD EG
(QUO ‘yprAcug) seus ‘wpovevgs sudasy T| (Dg ‘xe’ woIss) spews ‘wyavungo sudasy |g (GV “ANY ASOT) spew ‘WpvLvG9 sudasy OG (VA\ “OD eUIPyeA) speur ‘Wov«vGa
SUPSY 6% (OD “OD Wusuns) seu ‘wpovLvY9 sUdasy gH (YD “OD O82Iq UeS) ae ‘wpvLvGI sUdasy Lp (YD “OD O82Iq] ues) aeuI ‘wsounfiad sudasy Qy (VD “OD
o8aiq] ues) speuray ‘vsownfiad sudasy gp (YD “OD O821C UeS) seUIAy ‘Vsounf sudasy PY (YD “OD O82Iq UKs) seu ‘vsounf sudasy EY ‘synpe sudasy *gg—EpY Seansi4

oN
MN

Revision of the genus Aseptis McDunnough...

‘puagoyopnasd
sudasy gg vuayoy sudasy yg vouvunf sudasy ¢Q vaussnssaf sudasy TQ vuln sudasy |g votugia sudasy QQ vjoaun{ sudasy 6 “eITeITUAS a[eWI s77dasy *GQ—6g SA4N3I4

80 Tomas Mustelin & Lars G. Crabo / ZooKeys 527: 57-102 (2015)

vesica is average in size for the genus with a single long apical cornutus and two small di-
verticula located at the base and mid-portion. The female bursa is 1.3x as long as wide,
has distinct signa, and a short slightly bent appendix bursae with a crenulate shape.

Most specimens of A. binotata can be recognized, despite the variation in this spe-
cies, by their brownish color and pale postreniform patch. They are most likely to be
confused with Paraseptis adnixa, which occurs with it in the Pacific Coast states. The
basal dash of A. binotata is short, not reaching the antemedial line, whereas that of
P. adnixa extends to the antemedial line. Structurally, the male vesica of P. adnixa is
coiled and lacks a large apical cornutus, and the female ductus bursae has a sclerotized
plate in its posterior wall, unlike those of Asepztis.

Distribution and biology. This species is widespread in western North America
west of south-central Alberta, Wyoming, and Nebraska. Along the Pacific Coast it oc-
curs from northern Mexico to south-central British Columbia. It can be found from
sea level to altitudes over 2000 m in a variety of habitats from dense forest to shrub
desert. In the south the flight begins in March at low elevations and in April to July
in the mountains. In the north the flight begins later and lasts into August. The larva
is green with a white and red lateral stripe. It feeds on numerous woody shrubs, in-
cluding Ribes spp. (Grossulariaceae), Oemleria cerasiformis (Torr. & Gray ex Hook.
& Arn.) (Rosaceae), Salix spp. (Salicaceae), Acer spp. (Aceraceae), and Symphoricar-
pos spp. (Caprifoliaceae) (Miller and Hammond 2000, Miller and Hammond 2003).
Hampson (1908) mentions Adenostoma fasciculatum Hook. & Arn. (Rosaceae) as a
foodplant and Crabtree and Leuschner (2000) found larvae on Prunus emarginata
(Douglas ex Hook.) D. Dietr., P. virginiana (L.) var. demissa (Nutt.) Torr., and P.
subcordata Benth. (Rosaceae).

Discussion. Despite the superficial variability of A. binotata the male and female
genitalia and CO1 barcodes of this species are remarkably uniform. Barcodes from
parts of its northern distribution differ by circa 1% from other populations but there
is almost no variation elsewhere, including the reddish forms in the Southwest or the
variable California populations. We do not advocate the use of subspecies in A. bino-
tata because the variation is clinal and because of significant variation in color within
populations.

The species name genitrix has often been misspelled as “genetrix”.

Aseptis catalina (Smith, 1899)
Figs 35, 36, 67, 84

Hadena catalina Smith, 1899: 261.

Type material. Hadena catalina: Lectotype [USNM, examined]. Type locality: Cat-
alina Springs, Arizona.

Diagnosis. An average or slightly smaller than average Aseptis species, wingspan
31.840.9 mm (n=25; range 30-33 mm), with a powdery pale yellow-tan forewing

Revision of the genus Aseptis McDunnough.... 81

with patchy contrasting darker gray markings including the filling of the reniform spot
and the adjacent medial area. The postreniform patch is large but only slightly lighter
in color than the fold portion of the medial area and the filling of the lines. The basal
and postmedial areas are darker. The reniform spot is large and it and the claviform
spot are filled with dark gray. The antemedial and postmedial lines are black, filled
with pale cream and the postmedial line is often followed by black and white dots
on the veins. The subterminal area is pale cream, the terminal line is a series of black
spots and the fringe is checkered. There is some variation in the color and tone of the
ground color, which can be very pale cream or more tan colored. Although the pattern
is complete, the maculation is usually indistinct.

The male genitalia are similar to those of A. binotata but the valve lacks the digitus,
the ampulla of the clasper is thicker and is oriented parallel to the dorsal valve margin,
and the dorsal apex of the cucullus is pointed slightly. The aedeagus and vesica are like
those of A. binotata except for the presence of a granulose area on the ventral apex of
the aedeagus. In the female, the corpus bursae is oblong, circa 2.4x as long as wide,
with a curved appendix bursae of nearly the same size. The papillae anales are unique
in the genus in that they are covered by sparse short needle-like setae with brush-like
very dense basal setae condensed into patches on the ventral sides.

Aseptis catalina can be recognized by its patchy pale-tan and gray forewing and is
unlikely to be confused with other Aseptis; however, they resemble superficially Tride-
pia nova (Smith) and Scotogramma densa Smith, both in the Hadenini, and are often
mixed with them in collections. It is easily separated from them by the eyes, naked
in Aseptis, but covered in fine hairs in the two hadenines, as well as by the lack of a
notched hindwing in these species.

Distribution and biology. ‘This species occurs in deserts of Arizona, California
and Baja California, Mexico. Most specimens are from the western edge of the Colo-
rado Desert in San Diego, Imperial, and Riverside counties, California, but there are
colonies throughout the Colorado, Mojave, and Sonora deserts. Like many desert in-
sects, the flight period depends on winter rainfall and is early, generally early March to
April. The food plants and immature stages are unknown.

Aseptis serrula (Barnes & McDunnough, 1918)
Figs 37, 38, 68, 85

Trachea serrula Barnes & McDunnough, 1918: 104.

Type material. Holotype male [USNM, examined]. Type locality: Palm Springs, Riv-
erside County, California.

Diagnosis. This below-average-sized Aseptis, wingspan 31.741.2 mm (n=19; range
29-34), is the only one with a serrate male antenna. The forewing is relatively narrow, pow-
dery gray, with the pointed black claviform spot as the most prominent mark. ‘The dark re-
niform and orbicular spots are less prominent, the basal, antemedial, and postmedial lines

82 Tomas Mustelin & Lars G. Crabo / ZooKeys 527: 57-102 (2015)

ad rs

Figures 66-75. Aseptis, Paraseptis, and Viridiseptis male genitalia. 66 Aseptis binotata 67 Aseptis catalina
68 Aseptis serrula 69 Aseptis torreyana 10 Aseptis susquesa 1\ Aseptis fumosa 72 Aseptis perfumosa 73 Aseptis
characta 14 Paraseptis adnixa 15 Viridiseptis marina.

are faint or absent, and the subterminal line is often evident as a pale W-mark on veins M3
and CuA1. The postreniform patch is relatively small, and the medial area is often lighter
than the ground color near the claviform spot. The hindwing is off-white with dark veins
and terminal area in males and darker gray with light base and dark veins in females.

Revision of the genus Aseptis McDunnough... 83

87 88

Figures 76-91. Aseptis, Paraseptis, and Viridiseptis female genitalia. 76 Aseptis fumeola 77 Aseptis ethnica
78 Aseptis murina 19 Aseptis ferruginea 80 Aseptis fanatica 8\ Aseptis lichena 82 Aseptis pseudolichena
83 Aseptis binotata 84 Aseptis catalina 85 Aseptis serrula 86 Aseptis susquesa 81 Aseptis fumosa 88 Aseptis
perfumosa 89 Aseptis characta 90 Paraseptis adnixa 9\ Viridiseptis marina.

84 Tomas Mustelin & Lars G. Crabo / ZooKeys 527: 57-102 (2015)

The male uncus is unique in that the subbasal segment is expanded and dorsoven-
trally flattened to an elongate rhomboid shape with a slight constriction at the end of
the swollen segment. The valve is most similar to those of A. catalina and A. torreyana,
with a gently-curving S-shape, a curved ampulla of the clasper that is directed distally,
a narrow pointed digitus, and a slightly pointed cucullus. The vesica is like that of A.
binotata. In the female, the papillae anales are unique in being covered dorsally by
short scales with a rugose medial margin. The bursa copulatrix is nearly round with
strong signa and the appendix bursae is as long as the corpus bursae and curved left-
ward and dorsad from its origin.

Males of A. serrula are distinguished easily by the serrate antenna, filiform in other
Aseptis. Females are readily identified by their unique papillae anales. Many gray desert
noctuids resemble A. serrula, including several species of Hadenini with which it is often
confused. Differences between Aseptis and hadenines are described under A. catalina.

Distribution and biology. This is a species of the lower mountain-desert transi-
tion zone and high desert and has been collected in the Mojave, Colorado, and Sonora
deserts of southeastern California, Nevada, Arizona, and Baja California. It flies during
the desert spring, between March and early May depending on winter rainfall. Its host
plants and immature stages are unknown.

Aseptis torreyana Mustelin, 2006
Figs 39, 69

Aseptis torreyana Mustelin, 2006: 29.

Type material. Holotype male [SDNHM, examined]. Type locality: Torrey Pines
State Reserve, La Jolla, California.

Diagnosis. This is the smallest and rarest Aseptis (wingspan 27.5 mm). It has a
pale buff forewing overlaid with pale gray scales and marked with black basal dash
and outlines of the three spots. The reniform spot is the largest and is filled with dark
scales. The pale postmedial line is barely visible while the antemedial line is missing.
The strong black basal dash gives the impression of a small pale P. adnixa.

Males of A. torreyana have a narrow S-shaped valve with an ampulla of the clasper
that is oriented toward the apex of the valve, a small digitus that arises near the ventral
attachment of the clasper and is oriented perpendicular to the valve, and a small rounded
cucullus. The aedeagus and vesica resemble those of A. binotata. The female is unknown.

This species is unlikely to be confused with any other Aseptis because of its small
size and isolated habitat. It resembles a pale A. serrula, although the markings of A.
torreyana are more distinct. The male antenna of A. sorreyana is filiform rather than
serrate. Aseptis torreyana is also reminiscent of Aseptis characta (Grote) but is easily dis-
tinguished from it by the present of a digitus on the male valve.

Distribution and biology. This species is known only from the south side of the
sea level salt marsh estuary of the Torrey Pines State Reserve. The habitat is most likely

Revision of the genus Aseptis McDunnough.... 85

salt marsh, although it could be coastal chaparral. The capture date of April 21 is in
line with the spring flight period of most Aseptis species in southern California. The
foodplant is unknown.

Discussion. Aseptis torreyana is enigmatic because of its rarity. It was discovered
in a small isolated coastal chaparral remnant bordering the salt marsh within Torrey
Pines State Reserve, San Diego, California. Only two males were ever found, both at
this locality on the same night, despite a two-year moth survey by Norris Bloomfield
and the senior author. The latter also ran a blacklight nearly every night for five years
at the rim of the Pefasquitos Canyon, which runs into the same estuary 5 miles east
of the type locality without finding additional specimens. Similarly, none were found
at the nearby Miramar Air Station that was surveyed extensively for Lepidoptera for
years with same traps (Brown and Bash 2000). Taken together, it appears that Aseptis
torreyana is very local, perhaps tied to a food plant in the salt marsh. Another example
of such a restricted species is Orthomoia bloomfieldi Mustelin that was described from
30 specimens from a single locality during the Miramar study.

Aseptis susquesa (Smith, 1908)
Figs 40-42, 70, 86

Hadena susquesa Smith, 1908: 116.
Trachea monica Barnes & McDunnough, 1918: 104, syn. n.

Type material. Hadena susquesa: Lectotype male designated by Todd (1982) [AMNH,
examined]. Type locality: Claremont, California. Trachea monica: Holotype male
[USNM, examined]

Diagnosis. This is a slender-winged small to medium-sized Aseptis, wingspan
31.4+1.2 mm (n=25). It is readily recognized by its streaky medium-gray to dark-
brown gray forewing with streaks of warm light orange tan to yellow tan at the pos-
treniform patch, in the fold, and in the large pointed claviform spot. A thin tan line
parallels the margin near the anal angle. The reniform and orbicular spots are outlined
in black with paler peripheral and darker central scaling. The distal forewing is streaky
due to black veins and pale-gray scales abutting R5, M1, M3, and CuA1. The trans-
verse lines are obsolete. The hindwing is light whitish gray with brown-gray marginal
shading and dark veins, darker in females. Aseptis susquesa from coastal California tend
to be rustier than those from inland locations.

The male valve of A. susquesa is similar to that of A. binotata, although the cucullus is
larger. The uncus is thin, the valve has an upright ampulla, the oblique digitus is long and
pointed, and the cucullus is approximately 1.5x as wide as the valve and slightly pointed.
The vesica has two small diverticula and a single apical cornutus. The female genitalia
has a papilla analis covered by similar-length needle-like setae and sparse hair-like basal
setae; the corpus bursae is relatively small and short, 1.25x as long as wide, with a similar
sized appendix bursae that is laterally compressed and asymmetrically bulging ventrally.

86 Tomas Mustelin & Lars G. Crabo / ZooKeys 527: 57-102 (2015)

No other Asepztis is streaked gray with patches of light orange or rusty color. Aseptis
susquesa is most similar to A. serrula and flies with it. It is similar gray but has light
color restricted to a small postreniform patch. Males of these species are easily distin-
guished by their antennae

Distribution and biology. Aseptis susquesa is known from Arizona, California,
and Baja California, Mexico, at least as far south as Ensenada. Most records of the
rusty coastal form are from San Diego, Riverside, and Los Angeles counties, Califor-
nia, where it inhabits coastal chaparral and canyons from late March to early June.
The grayer inland form is found in the Mojave and Colorado deserts of southern
California and in the Sonora Desert of Arizona. Aseptis susquesa prefers rocky areas in
the mountain-desert transition zone and high desert. The larva is dark green marked
with white and feeds on Artemisia californica Less. (Asteraceae) (unpublished) and
Ericameria laricifolia (A. Gray) Shinners (Asteraceae) (Crumb 1956).

Discussion. In their original description of Trachea monica from Redington, Ari-
zona, Barnes and McDunnough (1918) pointed out that it is similar to Hadena susque-
sa, described previously from Claremont, California, and might be a gray inland form
of it. In support of that notion, the lectotype of Hadena susquesa is rather gray whereas
some desert specimens from farther inland have considerable rusty brown demonstrat-
ing variability and overlap. The male genitalia of these forms are virtually uniform
throughout the range. The CO1 barcodes, including specimens typical of coastal and
inland forms, vary by less than 0.8%. For these reasons, we treat Trachea monica Barnes
& McDunnough as a junior subjective synonym of Hadena susquesa Smith.

Aseptis fumosa (Grote, 1879)
Figs 43, 44, 71, 87

Hadena fumosa Grote, 1879: 205.

Type material. Holotype female [BMNH, photograph examined]. Type Locality:
Colorado.

Diagnosis. A medium-sized Aseptis, wingspan 34.9+1.5 mm (n=25; range 31.5—
38 mm) with a very dark forewing and whitish hindwing with black veins in males and
darker gray hindwing in females. The forewing is uniform smoky dark blackish brown
with brown filling of the antemedial and postmedial lines that is most evident as dots
on the costa. The weakly figure-eight shaped reniform spot, orbicular spot, and short
claviform spot are black filled with ground color or slightly darker scales. The male
hindwing is pearly gray distal to the spot that accentuates the vein asymmetry. The
hindwing of the female is smoky dark with dark but less conspicuous veins.

The male genitalia of A. fumosa are unique in several respects. The valve extends
nearly 90° lateral from its support and is a simple strap with a narrow base, very weak
sacculus, convex ventral mid-portion, and undifferentiated cucullus; the clasper is near
the base with a short straight ampulla parallel to the costa, and the digitus is absent.

Revision of the genus Aseptis McDunnough.... 87

The uncus is thin and cylindrical. The penicillus is weak with a pointed dorsal margin.
The vesica is similar to that of A. binotata but bears an additional thick-based thorn-
like mesial cornutus.

The female has a papilla analis covered with short needle-like setae and sparse basal
hairs. The corpus bursae is fairly short, 1.5x as long as wide, with strong signa and the
appendix bursae is box shaped and laterally compressed.

Aseptis fumosa males are distinctive due to the combination of blackish forewing
and white hindwing with dark veins. Despite this, A. fumosa is often confused with
A. perfumosa in collections. Females of both species have dark hindwings and are less
easily separated. Aseptis fumosa is usually larger with a broader forewing and its spots,
especially the claviform spot, are less prominent than in A. perfumosa. Aseptis fumosa
females can also be confused with dark species in the Aseptis fumeola species group
such as A. ethnica and A. fanatica. The shapes of the bursae are distinctive. The blunt
rounded appendix bursae of A. fumosa distinguishes it from all of the look-alikes.

Distribution and biology. Aseptis fumosa is widespread in western North America
and is known from western Canada, Washington, Nebraska, Colorado, Utah, Arizona,
Nevada and California. It occurs in a variety of diverse habitats including coast chapar-
ral, dry conifer forest, and shrub steppe; it is not found in mesic forests. The flight be-
gins in April or May and lasts to July. The larva is smooth green with a white subdorsal
stripe and broad red and white lateral stripe (Miller and Hammond 2003). It has been
reared on Purshia tridentata (Pursh) DC., Cercocarpus sp. and Adenostoma fasciculatum
Hook. & Arn. (all Rosaceae) (Crumb 1956, Crabo et al. 2012). At higher altitudes in
the Cascades and in southwestern Oregon it feeds on Ceanothus integerrimus Hook. &
Arn. (Rhamnaceae) (Miller and Hammond 2000).

Discussion. The simple valve of A. fumosa is similar to that of A. characta. Both
of them also have multiple cornuti on the vesica. Although these derived states of the
valve suggest a close relationship, their female bursae differ in shape and that of A.
characta lacks signa.

Aseptis perfumosa (Hampson)
Figs 45, 46, 72, 88

Trachea perfumosa Hampson, 1918: 131.

Taxonomy. The type material of Trachea perfumosa Hampson, the holotype female
and two paratype females, was originally part of the type series of Trachea fumeola
Hampson 1908. In fact, the female later selected as the 7: perfumosa holotype was
depicted on plate 112 as T. fumeola. The male and females were described in separate
paragraphs and the male was selected as the type of 7. fumeola. Hampson later realized
that these females and the male type were separate species, naming the females Trachea
perfumosa Hampson, 1918. The 1918 work lacks a description—initially leading us to
suspect that Trachea perfumosa is a nomen nudum—but instead references the female

88 Tomas Mustelin & Lars G. Crabo / ZooKeys 527: 57-102 (2015)

T. fumeola description and illustration in the earlier work. This indication thereby
validates the name.

Type material. Holotype: female [BMNH, photograph examined]. Type locality:
USA, California.

Diagnosis. This is a small dark Aseptis with a wingspan of 32.7+1.2 mm (n=25;
range 30.5-34.5 mm). The body appears short and stout and the wings short and
stubby. The forewing is slightly mottled dark gray brown, almost black in some speci-
mens, often with a few grayish, brownish, olive, or reddish scales in the medial area
around the velvety black spots of which the acute claviform spot is usually the most
prominent. Less conspicuous forewing markings include a short black basal dash, in-
complete faint wavy basal, antemedial and postmedial lines filled with brown, and
irregular complete brown subterminal line. The hindwing of both sexes is dark grayish
brown with inconspicuous veins.

Males of A. perfumosa are separated easily from all other species of Aseptis by the
short spike-like digitus that arises near the ventral margin and is perpendicular to it.
The 90° basal bend of the vesica and basally-constricted medial diverticulum are also
diagnostic. Females are identified by the thin flange on the tip of each papilla analis
and by the bilobed appendix bursae.

Superficially, A. perfumosa is most similar to A. fumosa. Aseptis perfumosa tends to
be smaller and narrower winged than A. fumosa. Males are separated easily by hind-
wing color, dark in A. perfumosa and pale in A. fumosa. Females can often be separated
based the maculation as the claviform spot is usually the most prominent feature on A.
perfumosa, whereas the postmedial line is clearer in. A. fumosa. The correct identity can
be confirmed by examining the tips of the ovipositors under magnification.

Distribution and biology. Aseptis perfumosa is endemic to southern California
where it occurs in many habitats such as coastal chaparral and canyons, urban areas,
brush land, and open oak forest from sea level to 2000 m. It is often very common
and can be the most abundant noctuid species. The peak of its flight is early April to
early June in coastal areas and a little later at higher elevations. The larva and pupa are
described and figured by Comstock (1940b). The green larvae were found and reared
on Manzanita (Arctostaphylos spp.).

Discussion. Hampson (1918) gives the expanse of the type of Trachea perfumosa
as 36 mm. This is larger than any A. perfumosa specimens examined by us. This led
us initially to question whether he might have described a female of the larger species
A. fumeola rather than A. perfumosa, given that the latter species was named from a
specimen in the type series of the former species. While enquiring about the types at
the BMNH, Alberto Zilli (pers. comm. 2015) explained that Hampson’s wingspans
are almost always greater than those of the actual moth because of his method of
measurement. He measured from the pin to the apex of the forewing and doubled the
result. While the actual wingspan of the female type of Trachea perfumosa is 33.5 mm,
a normal size for the species, the result using Hampson’s method yields the published
result of 36 mm.

Revision of the genus Aseptis McDunnough.... 89

Aseptis characta (Grote, 1880)
Figs 47-52, 73, 89

Hadena characta Grote, 1880: 243.

Hadena luteocinerea Smith, 1900: 468.
Hadena erica Smith, 1905: 258.

Hadena pluraloides McDunnough, 1922: 237.

Type material. Hadena characta: Holotype female [BMNH, photograph examined].
Type locality: Nevada. Hadena luteocinerea: Holotype male [USNM, examined].
Type Locality: Montana. Hadena erica: Lectotype male designated by Todd (1982)
[AMNH, examined]. Type Locality: Stockton, Utah. Hadena pluraloides: Holotype
female [CNC, examined]. Type Locality: Lethbridge, Alberta.

Diagnosis. This is a narrow-winged smaller Aseptis, wingspan 32.0+1.2 mm
(n=25; range 29.5-35 mm), with complete forewing pattern of typical lines and spots.
The forewing is ash gray to pale tan, often darker gray in the medial area and with vari-
able olive-gray, tan, or orange-tan patches. The basal, antemedial and postmedial lines
are double, dark gray filled with pale gray. The postmedial area is lighter with a shade
preceding the pale subterminal line and a number of black wedges between the veins,
The three spots are outlined in black and are filled with the ground color and, except
the claviform spot, peripheral lighter gray scales. The hindwing is gray, pale gray, or
white with dark discal spot, veins, and terminal line, darker in females.

Aseptis characta is geographically variable, appearing slightly different in each re-
gion. In extreme southern California it is relatively smooth gray with a white hindwing
(Fig. 47), becoming more powdery with a gray hindwing in Los Angeles and San Ber-
nardino counties and more mottled in the Sierra Nevada. In the Pacific Northwest it
is usually darker gray with variable subtle olive or brown shades on the forewing and a
fuscous hindwing (Figs 49 and 51), although populations from the Blue Mountains of
Washington and Oregon resemble those from the Sierra Nevada. On the Great Plains
A. characta is lighter, often pale tan with a warm orange cast (Fig. 50).

The male valve is strap-like with a slight S-shape, with a very weak sacculus, slightly
expanded cucullus with a rounded apex, rod-like straight ampulla oriented parallel to
the costa, and no digitus. The vesica is similar to that of A. binotata but has one or
two additional spine-like cornuti on its mid-portion. The female has a papilla analis
covered densely with short needle-like setae and sparse basal hairs, a rounded corpus
bursae lacking signa, and a short appendix bursae that barely changes the outline of
the bursa.

Aseptis characta can be identified by its small size, mottled gray forewing, and com-
plete pattern of lines and spots. The male is the only Asepzis with two or three slender
cornuti on the vesica and the female is the only one with an immaculate corpus bur-
sae and weak appendix bursae. This species is similar to several species of Lacinipolia
McDunnough in the Eriopygini, especially Lacinipolia pensilis (Grote), and is often

90 Tomas Mustelin & Lars G. Crabo / ZooKeys 527: 57-102 (2015)

intermixed with them in collections. They can be distinguished by the hindwing notch
of Aseptis and minute hairs on the eyes of Lacinopolia.

Distribution and biology. Aseptis characta is widespread in western North Amer-
ica in the western Great Plains, Great Basin, and Pacific regions from British Colum-
bia, Alberta, and Saskatchewan to Colorado, Utah, northern Arizona and southern
California. It does not occur on the immediate Pacific Coast north of central Califor-
nia. It flies in dry habitats like sagebrush steppe, juniper woodlands, and open forest
from sea level to 2500 m and is often common. In southern California it is most often
found on the dry side of the mountain ranges, in the mountain-desert transition zone,
and in the deserts. Emergence is earliest in xeric habitats, usually April in California
and mid-May in the Pacific Northwest. The flight lasts until July to August depending
on locality. The striped gray-green and white larva has been found feeding on Artemisia
spp. (Asteraceae) (Comstock 1955, Crumb 1956).

Discussion. Similarities between the male genitalia of this species and A. fumosa
are noted under the latter species. The female corpus bursae lacking signa and shallow
appendix bursae of A. characta are unique.

Given the geographic variability of A. characta it is almost surprising that not more
names have been given to the various forms. Hadena erica Smith was based on speci-
mens from Utah which are bluish ash gray with patches of paler gray and an ochreous
tinge on the basal and distal wing similar to Fig. 48. The light orange-tan Great Plains
populations were described twice, as Hadena luteocinerea Smith from Montana and
Hadena pluraloides McDunnough from Alberta; both are similar to Fig. 50. The latter
name denotes the resemblance to Euxoa pluralis (Grote). Aseptis characta has a nearly
continuous distribution within its range and the different forms are not well enough
separated to warrant the use of subspecies.

In contrast to the variation in habitus of this species, the genitalia are uniform.
Similarly, the variation of CO1 barcode sequences is small despite a large number of
samples (n=67) from throughout its distribution. Multiple slightly different haplo-

types cluster within a total range of less than 1%.

Genus Paraseptis gen. n.

http://zoobank.org/E3418CA4-70AB-4F26-8EE6-ED770970B171

Type species. Hadena adnixa Grote.

Etymology. The name Paraseptis is derived from para meaning next to and septis
by analogy to Aseptis. The name is feminine.

Diagnosis. Paraseptis is a monotypic genus whose sole member occurs near the
Pacific Coast of North America. It is mottled brown with typical noctuid wing mark-
ings, including a basal dash and dark wedges near the outer margin, and a pale off-
white to ochre postreniform patch. The outer edge of the hindwing is concave focally
between M1 and M3 as in Aseptis and Viridiseptis. The male genitalia resemble those
of Aseptis but differ as follows: the valve has a much larger sacculus that extends above

Revision of the genus Aseptis McDunnough.... al

the costal margin, a twisted upright ampulla, a rod-like digitus arising near the base of
the cucullus from a longitudinal bar near the costa; a long curved aedeagus with distal
spine patches of small spines; and a coiled vesica with basal and medial cornuti, but
no long apical cornutus (occasional specimens with a minute apical cornutus). The
female genitalia are also similar to those of Asepzis but differ in having more narrow
papillae anales and a strongly sclerotized posterior ductus bursae. CO1 DNA barcodes
of Paraseptis are not similar to those of Aseptis and cluster variably with other genera
in the Xylenini when representatives from a large number of species are included in
the sample set.

Description. Adults: Head: Eye rounded, normal sized. Antenna filiform in both
sexes. Labial palpus unmodified with longer second segment and short third segment.
Frons slightly convex, smooth. Thorax: Paired moderate-sized dorsal mesothoracic and
smaller metathoracic tufts. Legs without tibial spines; tarsal segments with three rows
of short spine-like setae. Abdomen: Male with coremata at base of abdomen, complete
with lever, pocket, and Stobbe’s gland; proximal segments with weak dorsal tufts.
Forewing: Venation as typical for subfamily, approximately 0.6x as wide as long, with
brownish, black, and off-white scales, appearing mottled brown with black typical
noctuid markings including a basal dash. Hindwing: Venation trifine as typical for
subfamily, M2 weak but usually visible, clustered close to M3 and CuA1; outer mar-
gin contour concave between veins M1 and M3. Male genitalia (Fig. 74): Tegumen
narrow near base of uncus; penicillus large, quadrate. Uncus smoothly downcurved,
narrow, tapering smoothly from base to acute tip. Juxta rectangular, circa 2/3x as wide
as long. Valve weakly S-shaped, tapered from base to mid-portion then even in width
to base of cucullus; sacculus strong, moderately sclerotized, 0.4x valve length and 2x
valve width at base of ampulla, extending above costa; cucullus weakly constricted at
base and expanded to 1.5—1.7x valve width, slightly rounded with blunt apex, corona
of circa 30 claw-like setae; clasper on mesial third, ampulla oriented perpendicular to
valve and extending above costa, rod-like with slightly twist to mediolaterally flattened
tip; digitus at distal end of an evenly-thick sclerotized ridge located slightly below
costa from clasper to digitus origin near cucullus base, rod-like with blunt tip, ori-
ented 45° to valve, ending near ventral cucullus. Aedeagus narrower and more robustly
sclerotized than in Aseptis, 7x as long as wide, distal half bent ventrad approximately
60°, small patches of small spines near dorsal and ventral apex; vesica slightly wider
than aedeagus, coiled 360° to right and ventrad to end posterior, ventrad, and left of
aedeagus tip, with subbasal patch of short cornuti on right, single spike-like diver-
ticulum perpendicular to axis on anterior distal third, minute spike-like cornutus di-
rected basad at apex in a few specimens, and very small dome-like mesial diverticulum.
Female genitalia (Fig. 90): Papillae anales weakly sclerotized, asymmetrically cone-
shaped with point near dorsum, circa 1.7x as long as wide, covered posteriorly and
apically with short thin setae that are slightly shorter near tip, lacking hair-like basal
setae; apophyses moderately long, posterior apophysis 1.7x anterior apophysis; ostium
bursae membranous except for thin weak band in ventral wall; ductus bursae tubular,
1x corpus bursae length, proximal 2/3 sclerotized with longitudinal ridge in dorsum,

92 Tomas Mustelin & Lars G. Crabo / ZooKeys 527: 57-102 (2015)

distal 1/3 membranous; corpus bursae ovoid, 0.75x as wide as long, with four long
signa evenly spaced on anterior, posterior, and lateral sides; appendix bursae arising
from right paramedial ventral posterior corpus bursae, moderately sclerotized, rugose,
0.75x corpus bursae length, conical with 45—60° rightward bend to end ventral to, or
slightly to right and ventral to, distal ductus bursae, with ductus seminalis near apex.

Discussion. The structural differences of Paraseptis and Aseptis are surprising given
the nearly identical habitus of P. adnixa and A. binotata, which are often mixed in
collections.

Several similar features of Paraseptis and Aseptis, including the hindwing shape and
superficial resemblance, suggest that these genera are related closely. The hindwing
shape is rare in other genera in the Xylenini. It is a prominent feature of the Eurasian
monotypic genus Atypha Hiibner. The male genitalia of Atypha pulmonaris (Esper),
illustrated by Fibiger and Hacker (2007), are similar to those of A. fumosa and A. char-
acta in having a simple valve, horizontal ampulla, and no digitus, suggesting that these
three genera might share a common ancestor.

Paraseptis adnixa (Grote, 1880), comb. n.
Figs 53-56, 74, 90

Hadena adnixa Grote, 1880: 243.
Hadena pausis Smith, 1899: 262, syn. n.

Type material. Hadena adnixa: holotype male [BMNH, photograph examined]. Type
locality: Nevada. Hadena pausis: lectotype male designated by Todd (1982) [USNM,
examined]. Type locality: Los Angeles County, California.

Diagnosis. A medium-sized noctuid, wingspan 34.9+1.3 mm (n=25; range 32—
37.5 mm), that resembles strongly a narrow-winged A. binotata in color and pattern.
It is gray brown, has a full complement of dark lines and spots and a pale postreniform
patch, and has a streaky hindwing with dark veins. The basal dash is thicker black
than that of A. binotata and extends fully to the antemedial line. Black wedges on the
wing distal to the lower cell and in the fold are also more prominent in P. adnixa. In
the Pacific Northwest, P. adnixa is typically patchy brown, often with a reddish tint,
with a large pale ochre postreniform patch and contrasting black markings (Fig. 56).
In most of California, it tends to be paler and less well marked with a speckled gray-
brown or brown-gray forewing, less conspicuous postreniform patch and black marks,
and slightly lighter hindwing. This form is even more similar to A. binotata from the
same region (Figs 53, 54). In the vicinity of Mono Lake in east-central California, P.
adnixa is powdery pale gray with red-brown basal and postmedial areas, an off-white
postreniform patch, and more dark streaks on the distal wing (Fig. 55).

Differences in genitalia between Paraseptis and Aseptis are described under the Par-
aseptis genus description. Paraseptis adnixa can usually be identified without dissection

Revision of the genus Aseptis McDunnough.... 95

by the combination of notched hindwing, brown forewing with pale postreniform
patch, and long basal dash.

The CO1 barcodes of Paraseptis based on 59 samples from British Columbia to
southern California demonstrate six major haplotype clusters separated by at least
0.5% (Fig. 92). Of these, three clusters separated by at least 1.8% (PAD4, PADS,
PAD6) are from California west of the Sierra Nevada divide. The other two are more
divergent: two from southwestern British Columbia (PAD1+PAD3) and a single
Washington specimen, and the other from near Mono Lake, California (PAD2). The
BC/WA cluster differs by at least 2.6% from the Mono cluster and 2.5% from the CA
cluster, and the Mono and CA clusters differ by at least 2.8%. Interestingly, the three
most divergent clusters (BC/WA, Mono, and CA) correlate with the geographic vari-
ation described above. Although the alignment of barcode haplotypes and phenotypes
could suggest the presence of more than one species, no consistent male or female
genitalia differences were found to suggest the presence of more than a single species,
and all male genitalia match those of the lectotype of Hadena pausis at USNM (slide
#54). We therefore conclude that Hadena pausis Smith is a junior subjective synonym
of Hadena adnixa Grote.

Distribution and biology. This species is widely distributed along the Pacific
Coast from northern Mexico to southwestern British Columbia. It is found mostly
west of the divides of the Sierra Nevada, Cascades, and British Columbia Coast Moun-
tains but there are at least three colonies east of these mountains: Inyo and Mono
County, California; Klamath and Lake counties, Oregon; and interior British Colum-
bia near Lillooet. Paraseptis adnixa is common in the Pacific Northwest, where it can
be found in a variety of forested habitats. In southern California, it occurs in coastal
chaparral and in oak and brush land in the foothills and mountains. The flight period
is April to June in California and mid-May to August in the Pacific Northwest. The
larva feeds on Indian plum (Oecmleria cerasiformis) in the Rosaceae (Miller and Ham-
mond 2000) and might be a specialist on it in parts of its range (including the Pacific
Northwest). It has also been reported as feeding on Prunus (Rosaceae), which is the
likely foodplant where the moth is found outside of the range of Oemleria, such as in
interior British Columbia, south-central Oregon, and far-eastern California..

As described above, P. adnixa has three distinct populations based on superficial
appearance and CO1 barcodes. Although there is little evidence to suggest more than
a single species, we considered using subspecies to distinguish these forms. The Pacific
Northwest populations are continuous to the border with California (Crabo et al.
2012) suggesting that a cline to the California form may exist in northern California.
Until this is refuted it is best to consider these forms the ends of a north-south cline. By
contrast, the eastern California populations near the border with Nevada are probably
isolated. If distinguishing them with a subspecies epithet is desired the type locality
of Hadena adnixa Grote should be restricted since the stated type locality, Nevada,
could refer to Nevada or an unspecified site in eastern California (Lafontaine JD pers.
comm. 2015).

94 Tomas Mustelin & Lars G. Crabo / ZooKeys 527: 57-102 (2015)

1% perfumosa APE1 ay
| perfumosa APE2 f

perfumosa APES

fumosa AFUM be i ™
ferruginea AFE a cg
fanatica AFA1 i
" fanatica AFA2

ethnica AET1

ethnica AET2

ethnica AET3

murina AMU

fumeola AFUE1
fumeola AFUE2

susquesa ASU1 be cd
susquesa (‘monica’) ASU2
susquesa (‘monica’) ASU3 ax =f

, catalina ACA2

catalina ACA1 bid

catalina ACA3
serrula ASE

forreyana ATO

lichena ALI "sect?
pseudolichena APS hava

- binofata ABI2

binotata ABI1

binofata (‘genitrix') ABI6 Saye

binotata (genifrix’) ABIS =

binofafa (butata’) ABI3
binotata (‘dilara') ABI4
binofata (genitrix') ABI?
— characta ACH2
characta ACH4

characta ACH1
characta ACH3
Figure 92. Neighbor-joining CO1 tree of Aseptis. The letter and number code after each species is the

haplotype identifier as indicated in Table 1. An illustrative specimen for each species is shown on the right

hand side. For binotata and susquesa, the geographical forms are also shown.

Revision of the genus Aseptis McDunnough.... 95

Genus Viridiseptis gen. n.
http://zoobank.org/35396DDB-5784-4ACB-B5F9-821760B83BAF

Type species. Hadena marina Grote, 1874.

Etymology. The name is derived from viridis meaning green and septis by analogy
to Aseptis and Paraseptis. The name is feminine.

Diagnosis. Viridiseptis is a monotypic genus whose only member is found in Cali-
fornia and adjacent Oregon. It is a stout small to medium-sized moth with a powdery
mottled-green forewing.

The genitalia differ greatly from those of Aseptis and Paraseptis. In the male the
distal half of the uncus is broad, flat, and covered densely by short fine hairs; the valve
has triangular process from the inner surface of the sacculus, lacks an ampulla, has a
thick blunt digitus, and the distal end is rhomboid without a typical cucullus or co-
rona. The females of these genera are also highly divergent. That of Viridiseptis has soft,
pad-like ovipositor lobes, very short apophyses, a sclerotized plate in the ventral wall
of the proximal ductus bursae, and a membranous corpus bursae lacking signa with a
membranous appendix bursae arising perpendicularly from the wall of the posterior
corpus bursae rather than as a sclerotized posterior extension of it.

Description. Adult: Head: Antenna filiform in both sexes. Frons smooth. Eye round-
ed, normal size, naked. Labial palpus unmodified, with moderately long second segment
and short distal segment, reaching mid-eye. Thorax: Dorsal paired tufts on meso- and
metathorax. Legs with tibiae lacking spiniform setae; tarsal segments with three rows of
short spiniform setae on each segment. Forewing: Short and rounded, outer margin weak-
ly scalloped, covered with olive-green, gray, black, and off-white scales, pattern of ordi-
nary transverse lines, orbicular and reniform spots, but lacking distinct claviform spot and
dashes. Hindwing: Venation typical of trifine noctuids with vein M2 weak, M2 closest
to M1; outer margin contour weakly concave between veins M1 and M3, less prominent
than in Aseptis. Abdomen: Base of male with paired hair-pencils, complete with levers and
pockets. Weak dorsal scale tufts on proximal segments. Male genitalia (Fig. 75): Tegumen
shape unmodified without narrower area near uncus; penicillus broad, rounded. Uncus
base, narrow, cylindrical, distal two-thirds thicker and wider, roughly canoe shaped with
proximal and distal tapered areas separated by even-width segment, dorsal distal portion
and undersurface of tip covered densely with short hairs. Saccus of vinculum long and
narrow. Juxta base broadly shield shaped, tapering to slight waist at junction with apical
third, apical segment at base of aedeagus expanded to slightly wider than “waist” with raised
sclerotized structure with slightly overhanging lateral edges and rounded tip. Valve narrow,
6x as long as narrow mesial section at end of sacculus, slightly curved dorsally; sacculus
0.4x as long as valve, reaching 2/3x to base of costa, with tooth-like triangular process on
distal portion near ventral part of clasper; cucullus large, costal portion thick and apex and
ventral portions thin, rhomboid with three points: right-angle point at dorsal base, slightly
acute apex lacking a corona, and more rounded and obtuse ventral margin; clasper reduced
to attachment on valve, ampulla absent; digitus arising at ventral cucullus from weak plate
on ventral distal valve, stout, short, tooth-like or curved ventrad. Aedeagus 5x as long as

96 Tomas Mustelin & Lars G. Crabo / ZooKeys 527: 57-102 (2015)

wide, distal third curved slightly ventrad, with abrupt reduction in caliber at mid-point
from bull-nosed sclerotized ridge across ventral wall, a patch of distal striae with long exten-
sions onto vesica and patch of small spines near ventral apex; vesica slightly shorter than
aedeagus, bent nearly 90° ventrad and to right at base, then curved slightly leftward to end
with tip ventrad to end of aedeagus, with moderate-sized basally-constricted diverticulum
on posterior wall at 1/3 from base and smaller dome-shaped diverticulum on left wall at 2/3
from base, cornuti absent. Female genitalia (Fig. 91): Papilla analis broadly triangular with
rounded tip, covered by hair-like posteriorly-directed setae that are shorter and more dense
at tip; segment VIII and apophyses very short; median posterior 7" sternite concave at os-
tium bursae; ostium bursae broad, weakly sclerotized; ductus bursae cylindrical, 4x length
of segment VIII, membranous with granulose sclerotized plate with thicker posterior por-
tion forming a slight lip in ventral wall “4 distance from ostium to corpus bursae; corpus
bursae membranous without signa, pear shaped with narrow posterior and larger ovoid
anterior portions, ~1.7x as long as ductus bursae; appendix bursae extending ventrad and
slightly rightward perpendicular to corpus bursae from origin on posterior ventral wall of
corpus bursae, membranous, ovoid, ~1/3 size of corpus bursae, with junction with ductus
seminalis at left posterior base near corpus bursae.

Discussion. The higher classification of this genus is enigmatic. Although Virid-
iseptis clusters with genera in the tribe Xylenini by CO1 barcodes and its hindwing
notch suggests an affinity to Aseptis and Paraseptis, absence other structural similarities
between them and the biology of its larva suggest that the recent association with Asep-
tis is incorrect. The distal male valve of Viridiseptis bears some resemblance to other
genera in the Xylenini such as Sunira Franclemont, but the valve differs greatly in
other respects such as absence of the ampulla. A long twisted ampulla is one of the de-
fining characters of the subtribe Xylenina (Fibiger and Lafontaine 2005). In addition,
the larva of Viridiseptis feeds externally on forbs rather than woody plants, a defining
character of the entire tribe Xylenini (op. cit.), further clouding the relationship. There
is little evidence that Viridiseptis is related closely to Oligia Hiibner where it had been
placed (Franclemont and Todd 1983) prior to the most recent check list (Lafontaine
and Schmidt 2010). For these reasons, the phylogenetic position of Viridiseptis in the
Noctuinae is uncertain. We suggest that it be placed in the Xylenini (incertae sedis)
section pending a more encompassing revision of the subfamily.

The ridge on the ventral male aedeagus and plate in the ventral wall of the female
ductus bursae might be engaged during copulation.

Viridiseptis marina (Grote, 1874), comb n.
Figs 758,47 5594

Hadena marina Grote, 1874a: 67.

Type material. Holotype female [BMNH, photograph examined]. Type locality:
California.

Revision of the genus Aseptis McDunnough.... oF

Diagnosis. A medium-sized or slightly smaller than average stout species, wing-
span 31.941.5 mm (n = 25; range 29.5-35 mm). The forewing is granular mossy
green, occasionally yellowish green, with mottled dark-gray to black and light-green
pattern that obscures all but the darkest parts of the lines and spots. The darkest areas
are a small patch at the base of the trailing edge of the wing, the cell and fold in the
medial area, a bar on the costa preceding the subterminal line, and terminal area op-
posite the reniform spot. The relatively small reniform spot and round orbicular spot
are filled with peripheral whitish and central green scales. ‘The basal, antemedial and
postmedial lines are double, black filled with green, and the subterminal line is green;
all are sinuous and appear incomplete. The fringe is checkered green and gray. The
hindwing is light brown gray with a darker border.

This species can usually be identified by superficial appearance alone. If in doubt,
the male genitalia described under the genus description are diagnostic.

Distribution and biology. Viridiseptis marina occurs throughout coastal California
and in southwestern Oregon as far north as Douglas County. It is widely distributed
in southern California, where it is often common. It is found in many habitats such as
coastal chaparral, mountain forest, mountain-desert transition zone, and occasionally in
the deserts from sea level to at least 2000 m. It flies from April to early July. The larva
feeds on forbs and has been recorded on Nemophila spp. and Pholistoma auritium (Lindl.)
Lilja (Hydrophyllaceae) and Lithophragma spp. (Saxifragaceae) (Robinson et al. 2010).

Discussion

This revision should help clear up confusion in the genus Aseptis by facilitating iden-
tification of the species—in no small part impeded prior to this study by more names
than actual species—and by refining the generic relationships of species included with
Aseptis in recent check lists. Of the genera treated herein, Aseptis and Paraseptis are
retained in the tribe Xylenini, subtribe Xylenina and Viridiseptis is transferred to Xy-
lenini (incertae sedis).

Although our work is based primarily on structural evidence, we also accessed
a large pre-existing data base of CO1 barcodes available at BOLD and tested our
theories against it during the course of this work. We observed, for the most part, a
good correlation between structural and molecular data. Lack of significant variation
in two variable widespread species, A. binotata and A. characta, supported pre-existing
synonomies in both species and the proposed new ones in A. binotata, and was reas-
suring evidence that no cryptic species were being overlooked. As recently reported for
a number of Canadian moths (Zahiri et al. 2014), we found additional instances of
morphologically and biologically well-defined species with no more than 1.5% differ-
ence in CO1 DNA sequence, e.g., within the Aseptis fumeola species group (Fig. 92),
as well as examples of structurally well-delineated species within which there is greater
diversity of CO1 DNA. The latter was most evident in Paraseptis adnixa which has
several major barcode haplotype clusters that differ by up to 2.8% (Fig. 93).

98 Tomas Mustelin & Lars G. Crabo / ZooKeys 527: 57-102 (2015)

Viridiseptis marina VMA

adnixa PAD?

adnixa PAD1

Paraseptis adnixa PADS

adnixa PADS

adnixa PAD4

adnixa PADS

Figure 93. Neighbor-joining CO1 tree of Paraseptis and Viridiseptis. The letter and number code after
each species is the haplotype identifier as indicated in Table 1. Illustrative specimens for Viridiseptis ma-
rina and the three geographical phenotypes of adnixa are shown on the right hand side.

Acknowledgements

We are grateful to J. Donald Lafontaine for photographs of type specimens, barcode
sequence similarity trees from BOLD, invaluable advice and comments on the manuscript,
to Jeremy deWaard, Ontario Biodiversity Institute for the loan of specimens, to Thomas
E. Dimock for numerous specimens, and the late Ronald H. Leuschner for advice,
specimens, original descriptions, and the recognition of putative new species of Aseptis.

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Revision of the genus Aseptis McDunnough.... 101

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102 Tomas Mustelin & Lars G. Crabo / ZooKeys 527: 57-102 (2015)

Check list of the species of Aseptis McDunnough, Paraseptis Mustelin &
Crabo, and Viridiseptis Mustelin & Crabo

Aseptis McDunnough, 1937
fumeola (Hampson, 1918)
= probata (Barnes & McDunnough, 1910)
ethnica (Smith, 1899)
murina Mustelin, 2000
ferruginea Mustelin, 2000
fanatica Mustelin, 2006
lichena (Barnes & McDunnough, 1912)
pseudolichena Mustelin & Leuschner, 2000
binotata (Walker, 1865)
= rubiginosa (Walker, 1865)
= extersa (Walker, 1865)
= paviae (Strecker, 1874)
= curvata (Grote, 1874)
= genitrix (Grote, 1878)
inconspicua (Smith, 1893), nomen nudum
= dilara (Strecker, 1899)
= bultata (Smith, 1906)
= cara (Barnes & McDunnough, 1918)
catalina (Smith, 1899)
serrula (Barnes & McDunnough, 1918)
torreyana Mustelin, 2006
susquesa (Smith, 1908)
= monica (Barnes & McDunnough, 1918)
fumosa (Grote, 1879)
perfumosa (Hampson, 1918)
characta (Grote, 1880)
= erica (Smith, 1905)
= [uteocinerea (Smith, 1900)
= pluraloides (McDunnough, 1922)
Paraseptis Mustelin & Crabo, 2015
adnixa (Grote, 1880)
= pausis (Smith, 1899)
Viridiseptis Mustelin & Crabo, 2015
marina (Grote, 1874)

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Full text of "Revision of the genus Aseptis McDunnough (Lepidoptera, Noctuidae, Noctuinae, Xylenini) with a description of two new genera, Paraseptis and Viridiseptis"

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